Salsola kali
(common saltwort)

Pictures

Identity

Preferred Scientific Name

• Salsola kali L.

Preferred Common Name

• common saltwort

Other Scientific Names

• Kali australis (R.Br.) Akhani & Roalson
• Kali tragus Scop.
• Salsola aptera Iljin
• Salsola australis R.Br.
• Salsola australis var. strobilifera (Benth.) Domin
• Salsola brachypteris Moq.
• Salsola kali f. hirsuta Hornem.
• Salsola kali subsp. austroafricana Aellen
• Salsola kali subsp. iberica (L.) Rilke
• Salsola kali subsp. kali
• Salsola kali subsp. pontica (Pall.) Mosyakin
• Salsola kali var. angustifolia Fenzl
• Salsola kali var. brachypteris (Moq.) Benth.
• Salsola kali var. glabra Forssk.
• Salsola kali var. hispida Forssk.
• Salsola kali var. hispida-polygama Forssk.
• Salsola kali var. kali
• Salsola kali var. leptophylla Benth.
• Salsola kali var. pontica Pall.
• Salsola macrophylla R. Br.
• Salsola pontica (Pall.) Iliin
• Salsola tragus L.
• Salsola tragus subsp. iberica Sennen & Pau
• Salsola tragus subsp. pontica (Pall.) Rilke
• Salsola tragus var. australis (R.Br.) Bég.
• Salsola turgida Dumort.

International Common Names

• English: prickly Russian thistle; prickly saltwort; Russian thistle; tumbleweed
• Spanish: barilla pinchosa; barrila borde; capitana; salicorn
• French: soude
• Portuguese: barrilha-espinosa; soda

Local Common Names

• Germany: Kali- Salzkraut
• Italy: erba cali
• Netherlands: Kalikruid; Loogkruid
• Sweden: sodaoert

EPPO code

• SASKA (Salsola kali)

Summary of Invasiveness

Many tumbleweeds (genus Salsola, section Kali) are important roadside, rangeland and agricultural weeds native to much of Europe, North Africa, the Middle East, Central Asia, China and Australia. Species have also been widely introduced accidentally as contaminants of crop and pasture seeds, and tumbleweed has become a widespread weed in Canada, USA, Mexico, Chile, China, Indonesia, Japan, Australia and New Zealand. S. kali and it many confused synonyms (including S. tragus and others which are disputed, such as S. iberica and S. australis) are common agricultural, rangeland and environmental weeds with multiple and important impacts. Major efforts have been made regarding mechanical, chemical and, more recently, biological control.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Plantae
•         Phylum: Spermatophyta
•             Subphylum: Angiospermae
•                 Class: Dicotyledonae
•                     Order: Caryophyllales
•                         Family: Chenopodiaceae
•                             Genus: Salsola
•                                 Species: Salsola kali

Notes on Taxonomy and Nomenclature

The genus Salsola is in the Chenopodiaceae, subfamily Salsoloideae, tribe: Salsoleae (USDA-ARS, 2015); however, some authorities place it in the Amaranthaceae family (The Plant List, 2013; Missouri Botanic Gardens, 2015a).

Three infraspecific taxa are accepted by The Plant List (2013): subsp. tragus (L.) Celak., subsp. ruthenica (Iljin) Soó and var. caroliniana (Walter) Nutt. However, USDA-ARS (2015) treats S. tragus as a separate species. Nonetheless, with no overall agreed view of specific limits, for the purposes of this datasheet the Plant List (2013) is taken as the definite opinion on taxonomy, with S. kali taken in its broadest sense. With S. tragus therefore accepted here as a synonym of S. kali, a number of further taxa are also drawn into to synonymity: S.tragus var. australis (R.Br.) Bég., S.tragus subsp. iberica Sennen & Pau, and S.tragus subsp. pontica (Pall.) Rilke.  However, where distribution or other records refer specifically to S. tragus, this will be noted as much as is practical, in the eventuality of any future revision of the genus that raises the tragus taxon to species level once again.

The Plant List (2013) also accepts S. australis and S. pontica as synonyms of S. kali, but accepts S. iberica as a separate species.

It is also important to include and note the naming authorities when considering synonymy. For example, USDA-ARS (2015) noted Salsola kali L. as separate from Salsola kali auct. non. (=Salsola tragus L.), whereas The Plant List (2013) recorded S. iberica Sennen & Pau as a good species, but S. iberica (Sennen & Pau) Botsch. ex Czerep as a synonym of S. kali subsp. tragus (L.) Celak.

In support of this, the following is the conclusion of the Flora of North American Editorial Committee (2015): 'Salsola tragus has been known in North American and European botanical literature under numerous names (for detailed synonymy see S. L. Mosyakin 1996 and S. Rilke 1999). Judging from the photographs of the Linnaean specimen of S. tragus (LINN 315.3), which should be regarded as a lectotype, it is the correct name for the widespread, narrow-leaved, weedy representative of the S. kali aggregate. In the present circumscription, Salsola tragus is an extremely polymorphic species consisting of several more or less distinct races (subspecies or segregate species). Several varieties may be recognized within S. tragus, many of them are just morphological variants of little or no taxonomic value. Studies using allozymes and DNA-based molecular markers in some North American and Eurasian representatives of Salsola tragus indicate that there are at least two cryptic genetically divergent populations (Ryan and Ayres, 2000). More studies may clarify distribution, origin, and taxonomic status of these infraspecific taxa (or cryptic species).'

S. kali is also referred to as Kali tragus (Akhani et al., 2007), based on a major reclassification of the genus that has not been generally accepted. Akhani et al. (2007) split the genus Salsola into five new genera, based on sections that were raised to the rank of genus. Although these genera are found in the literature, the broader limits of the genus Salsola is still generally accepted, which indicates some of the taxonomic confusion that still surrounds many Salsola species. The genus name Kali as proposed by Akhani et al. (2007; 2014) is not accepted here, as it appears to be not accepted by consensus taxonomical thought and understanding. The Plant List (2013) treats the genus Kali and all species therein either as synonyms or as ‘unresolved’.

Akhani et al. (2014) continued to argue that the correct generic name is Kali, describing the origins of the generic names Salsola L. and Kali Mill., and stating that the latter is legitimate name, published independently from Salsola, even though the original generic boundaries are by and large the same. This was countered, however, by Mosyakin et al. (2014), with a proposal to conserve the name Salsola (Chenopodiaceae s.str.; Amaranthaceae sensu APG), with Salsola kali as the conserved type.

Amongst species in Salsola sect. kali, commonly known collectively as the tumbleweeds, S. paulsenii is a relatively distinct species, unlike S. kali, S. tragus, S. australis and others that have a confused taxonomy, with numerous changes in synonomy and species limits. Regarding common names, Salsola species of section Kali are collectively know as the tumbleweeds. S. kali is commonly known as the Russian thistle, whereas S. tragus is referred to as the prickly Russian thistle, and S. paulsenii as the barbwire Russian thistle, giving some indication as to morphological differences by which the taxa are separated.

Beatley (1973) observed that Russian thistle (S. kali) populations widespread in western USA appeared to consist of two distinct species: S. paulsenii on disturbed land below 1200 m and especially on limestone-derived soils, and S. iberica abundant on disturbed land above 1800 m and especially on sandy soils of volcanic origin, although noting that hybridisation was frequent, with various degrees of introgression.

Nonetheless, Smith et al. (2013), presented a clear distinction between the exotic and invasive Salsola species present in North America, based on recent studies of morphology, allozymes and molecular genetics. These indicated that what is often knows as ‘Russian thistle’ comprises seven distinct species, with S. tragus probably the most widespread, S. collina Pall. mainly east of the Rocky Mountains, S. paulsenii Litv. primarily in deserts, S. kali restricted to ocean shores and is not a rangeland weed, and S. australis R. (sometimes known as ‘type B’), found mainly in California, South Africa and Australia, and has never been documented in Eurasia. Polyploid hybrids include S. x gobicola (including S. tragus and S. paulsenii) that is known in western USA and central Asia, and S. x ryanii (including S. tragus and S. australis) known only from California.

Although it was previously believed that all species in the kali Section of Salsola originated in Eurasia, the presence of four indigenous species in Australia suggests a separate clade, and it is likely that S. australis is native to Australia. However, other taxonomic listings such as the Plant List (2013), GRIN (USDA-ARS, 2015) and Tropicos (Missouri Botanic Gardens, 2015a) tend to include S. tragus, S. australis and at least some of their noted varieties as synonyms of S. kali.

Using alloenzymes and DNA-based molecular markers, Ryan and Ayres (2000) confirmed the distinction between two accessions of S. tragus [S. kali subsp. tragus] (one from California, USA, and the other from the native range of France and Turkey), and S. paulsenii. Analysis confirmed the genetic distinctness of the two isoenzymic phenotypes of S. tragus, and that S. paulsenii was markedly different from both types. Further analysis (Ayres et al., 2009) reconfirmed as distinct species S. tragus and S. paulsenii, as well as the recently identified S. kali subspecies austroafricana, possibly native to South Africa.

Molecular analyses also identified as an additional taxon a new allopolyploid hybrid between S. tragus and S. kali subsp. austroafricana, and another as a complex hybrid involving S. tragus, S. paulsenii and S. kali subsp. austroafricana.

Genetic variation was examined in S. tragus [S. kali subsp. tragus] from California (USA) using random amplified polymorphic DNA, intersimple sequence repeat analyses, and isoenzymes. A cryptic species, called type B, was found in addition to S. tragus. Type B has also been found in Arizona but not in other western states of USA. Examination of S. tragus from many locations in Europe, Asia, and Africa using DNA-based analyses has not revealed a source of type B.

Description

Adapted from the Flora of North America Editorial Committee (2015):

S. kali is a low herb, 5-50 cm tall, papillose to hispid or, occasionally, glabrous. Stems are erect to ascending, branched from base; with branches arcuate or occasionally almost prostrate. Leaves are alternate; blades linear, 1-2 mm wide, fleshy, usually not swollen at base, apex acuminate, forming a rather firm spine, 1-1.5(-2.2) mm long. Inflorescences interrupted at maturity, usually 1-flower per axil of bract; bracts alternate, not imbricate at maturity, reflexed, not distinctly swollen at base, apex narrowing into subulate spine. Flowers with bracteoles free or becoming connate and adnate to perianth base; perianth segments with comparatively narrow wing, or in lower flowers occasionally wingless, with weak or firm, acute apex, glabrous; fruiting perianth 4-6(-8) mm diameter.

The Flora of North America Editorial Committee (2015) also noted the following distinctions between the subspecies S. kali subsp. kali and S. kali subsp. pontica:

- S. kali subsp. kali: perianth segments having rigid, subspinose apex and prominent midvein; and bracteoles distinct, not swollen.

- S. kali subsp. pontica: perianth segments having weak apex and obscure midvein; bracteoles connate at base, swollen; flowers sometimes prominently winged.

In addition, the Flora of North America Editorial Committee (2015) provided a separate entry for S. tragus, including as synonyms: S. australis, S. iberica, S. kali var. tenuifolia Tausch and S. pestifer:

S. tragus is a low to medium herb, (5-)10-100 cm tall, sparsely papillose to hispid or glabrous. Stems are erect, rarely ascending, profusely branched from or near base (rarely simple in underdeveloped specimens); branches arcuate, proximal ones occasionally prostrate. Leaves are alternate; blades filiform or narrowly linear, less than 1 mm wide, not fleshy, not swollen at base, apex subspinescent, with spines less than 1.5 mm long. Inflorescences interrupted at maturity (at least proximally), 1-flowered (rarely 2-3-flowered with lateral flowers mostly abortive); bracts alternate at maturity, not imbricate, reflexed, not distinctly swollen at base, ± abruptly narrowing into mucronulate-spinose apex. Flowers with bracteoles distinct or occasionally connate at base in proximal flowers; perianth segments with prominent, membranous wing at maturity (two inner wings usually much smaller than the other three), apex weak, obtuse to weakly acuminate or reflexed, glabrous; fruiting perianth ca. 4-10 mm in diameter. 

Plant Type

Distribution

S. kali is reported as native to much of northern and western Europe, from Norway, Finland, the Baltic states and northern Russia, through Poland, Germany, the low countries, the UK, France, Spain and Portugal (USDA-ARS, 2015).                                                                                

S. kali is reported as introduced and naturalized in Canada, the USA, Mexico, Argentina, Chile, Australia and New Zealand (USDA-ARS, 2015). The Flora of North America Editorial Committee (2015) also records it as invasive in Central and South America and southern Africa.

As S. tragus (here accepted as a synonym of S. kali; see Notes on Taxonomy and Nomenclature), USDA-ARS (2015) records a broader native range than S. kali, encompassing that of S. kali in northern and western Europe, but also extending much further east and south, including the whole Mediterranean region, North Africa (including the Azores and Canaries), West Asia, the Middle East, Central Asia and Siberia, Pakistan, northern India, Nepal and China (USDA-ARS, 2015).

S. tragus is also widely reported as introduced. It is common to abundant in western and central parts of the USA, and found occasionally on eastern and southern coasts, where it is spreading rapidly (Duke, 1983), However, as stated in the Notes on Taxonomy and Nomenclature, uncertainty regarding nomenclature and species limits means that a more detailed review of the history of introduction, in light of a clearer taxonomic a breakdown, would provide valuable insights. In any case, S. kali/S. tragus is highly likely to be more widespread than indicated in the Distribution table.

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

S. tragus (here accepted as a synonym of S. kali; see Notes on Taxonomy and Nomenclature) was reported as being introduced to North America from Eurasia in the 1800s. It was probably introduced to South Dakota in 1870 or 1874 in flaxseed imported from Russia (Beatley, 1973). It is spreading rapidly on the eastern and southern coasts of the USA (Duke, 1983).

Habitat

S. kali appears to be well adapted to a very large range of habitats. Where naturalized, it is found along roadsides, disturbed areas, scrub vegetation, on former sugarcane fields, heath, shrubland and coastal areas (PIER, 2015). It is also a common on fallows, agricultural land and rangeland, especially abandoned fields and overgrazed pastures, shingle and sand behind beaches, and dry gravelly sites in New Zealand. As S. tragus, it is commonly occurs along roadsides and railways, and other dry, stony and sandy areas (Beckie and Francis, 2009).

Habitat List

Hosts/Species Affected

S. kali is common weed of a very wide range of agricultural crops, as well as grassland plants.

Biology and Ecology

Genetics

The chromosome number is reported to be 2n = 36, but this should be clarified, especially regarding the many synonymous taxa.

Gaskin et al. (2006) called the genus ‘taxonomically challenging’, and undertook morphological, cytological and molecular work to identify an unknown taxon, previously identified as S. tragus, but recently temporarily designated as Type B with unknown origins, present in California and Arizona. They found that it was morphologically very similar, and genetically identical, to S. kali subsp. austroafricana from southern Africa, and distinct from S. tragus. It is unclear if S. kali subsp. austroafricana is native to southern Africa or to the northern hemisphere of the Old World, and further investigations in both regions are needed (Gaskin et al., 2006).

Reproductive biology

The population dynamics of S. australis [S. kali subsp. tragus] were studied in its native range in Western Australia to facilitate the development of improved control strategies (Borger et al., 2009a,b). Two to three cohorts emerged in summer, and in autumn or winter, with an estimated seed production of approximately 100 to 20,000 seeds per plant. Seed viability was low, at 9% for shed seeds and 15% for seeds retained on mature plants. Total annual seed production per unit area was highly variable within sites but was not significantly different between sites (Borger et al., 2009a).

Borger et al. (2009a,b) detailed the taxonomy, morphology, geographical distribution, habitat, growth and development, reproduction, population dynamics, detrimental and beneficial attributes and management (chemical, physical and biological control) of S. australis. See also Crompton and Bassett (1985), discussing S. kali subsp. ruthenica as S. pestifer A. Nels.

Physiology and phenology

As a C4 species, S. tragus (here accepted as S. kali; see Notes on Taxonomy and Nomenclature) is highly competitive in semiarid and arid small-grain cropping systems because of its ability to emerge early, efficiently extract available water from the soil by its extensive root system, and to tolerate heat, drought and salinity. Moreover, the evolution of acetolactate synthase-inhibitor resistance has impacted herbicidal control of the species (Beckie and Francis, 2009).

Environmental requirements

S. kali is native to very wide range of cold, cool and temperate climates. Duke (1983) noted that S. kali ranged from cool temperate desert to steppe to subtropical very dry to thorn forest life zones. Correspondingly, it tolerates a very high range of annual rainfall and annual temperatures (9 to 24°C noted by Duke, 1983). However, subspecific taxa, subspecies, varieties, forms and ecotypes are very likely to be much more specific in their environmental preferences.

In its native range, this highly variable plant is common in coastal beaches and tolerates saline soils, but is also frequently found in inland areas. In the Himalayas, where it is native, it is found at least up to 4,500 m altitude (PIER, 2015).

Much more detailed analysis is required to elucidate specific environmental requirements following taxonomic clarification.

Soil Tolerances

Soil drainage

• free
• impeded

Soil reaction

• acid
• alkaline
• neutral

Soil texture

• light
• medium

Special soil tolerances

• infertile
• saline
• shallow
• sodic

Notes on Natural Enemies

Having been intensively investigated for possible biological control agents, there are many studies on the natural enemies of S. kali in its native range (see Biological control).

In 2005, dying S. tragus [S. kali subsp. tragus] plants were found along the Aegean Sea coast of Greece, with dark brown necrotic lesions on the stems and leaves caused by Colletotrichum gloeosporioides [Glomerella cingulata]. This is thought to be the first report of anthracnose on S. tragus in Greece (Berner et al., 2006). The same pathogen was found on the same species along the Azov Sea in Russia and was the first report of anthracnose caused by C. gloeosporioides on S. tragus in Russia (Kolomiets et al., 2008). In the same region, a further pathogen causing girdling, cracking and dark brown necrotic, canker-like lesions near stem bases, was identified as a Phomopsis, and the conidial state of Diaportheeres, thought to be the first report of stem canker on S. tragus caused by D. eres (Kolomiets et al., 2009).

Means of Movement and Dispersal

Natural dispersal

S. kali is a self-seeding annual, each plant producing up to a million seeds (Duke, 1983), and seeds may remain viable for 6-12 months. The dry, ball-shaped plant breaks off at ground level when mature and is rolled by the wind, spreading seeds as it goes (PIER, 2015). Pollen-mediated gene flow and efficient seed dispersal aids both short- and long-distance spread (Beckie and Francis, 2009).

Vector transmission (biotic)

S. kali is likely to be dispersed by small rodents who collect and store seed in caches, as is observed with other Salsola species. It is possibly also spread by birds, but further literature-based evidence is required to support this.

Accidental introduction

As S. tragus, it was reported as being introduced to North America from Eurasia in flaxseed imported from Russia (Crompton and Bassett, 1985). It likely that many other initial international introductions were accidental.

Pathway Causes

Pathway Vectors

Plant Trade

Impact Summary

Economic Impact

Tumbleweeds (genus Salsola, section Kali) are important roadside, rangeland and agricultural weeds in Australia, Chile, China, Indonesia, Japan, Mexico, New Zealand (PIER, 2015) and throughout the 48 contiguous states in the USA and the Canadian provinces (USDA-NRCS, 2015).

As a naturalized species, S. tragus (here accepted as S. kali; see Notes on Taxonomy and Nomenclature) is a common and economically important weed in crop production systems and non-cropped disturbed areas in semiarid to arid regions of western and eastern North America (Beckie and Francis, 2009), and a severe postharvest problem in the northwestern USA (Young et al., 2008).

Borger et al. (2009a,b) detailed detrimental and beneficial attributes of S. australis [S. kali subsp. tragus]. See also Crompton and Bassett (1985), discussing S. kali subsp. ruthenica as S. pestifer A. Nels.

Environmental Impact

Where invasive, S. kali displaces native plants and competes for space, water and nutrients (PIER, 2015). More work is required to elucidate the specific environmental impacts of S. kali and related species, such as its impact relative to native species diversity. For example, in Jasper National Park, S. tragus has invaded montane grasslands that provide critical winter range for bighorn sheep (Ovis canadensis Shaw) and other ungulates (Antill et al., 2012). S. tragus [Salsola kali subsp. tragus] was identified as a major concern on the Midriff Islands in the Central Gulf Coast region along the Sea of Cortés, off California (Bruckart et al., 2004).

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
• Pioneering in disturbed areas
• Highly mobile locally
• Fast growing
• Has high reproductive potential
• Has propagules that can remain viable for more than one year
• Has high genetic variability

• Monoculture formation
• Negatively impacts agriculture
• Reduced amenity values

• Competition - monopolizing resources

• Highly likely to be transported internationally accidentally
• Difficult to identify/detect as a commodity contaminant
• Difficult to identify/detect in the field
• Difficult/costly to control

Uses

S. kali and related species have been used as an emergency forage during drought. It is palatable when immature and non-toxic to livestock (Beckie and Francis, 2009). S. iberica and hybrids have also been used for making hay and silage in bad years with very few reports of ill effects (Fowler et al., 1979). S. kali is chemically comparable to many cultivated forages and appears amenable to cultural modification of its nutritive properties, and the considerable variability among intergrades between these species is an advantage here.

Uses List

Animal feed, fodder, forage

• Forage

Similarities to Other Species/Conditions

S. kali is similar to many Salsola species to the point where differences are obscure. Indeed, species limits are not agreed, resulting in confusing taxonomy. This is further complicated by the application of various names, especially where introduced, some of which are likely to have been misapplied.

S. kali is treated here as synonymous with S. tragus sensu stricto (see Notes on Taxonomy and Nomenclature), which is similar to S. paulsenii. Intermediate forms between species are known, and S. paulsenii × S. tragus hybrids have been identified, identical with S. × gobicola. Forms conventionally named as ‘Salsola X’ and ‘S. paulsenii lax form’ with 2n = 54 (e.g. Arnold, 1972; Ryan and Ayres, 2000) are probably also of hybrid origin. S. paulsenii is also noted to be similar in flowers and fruits to S. tragus and S. vermiculata, but it is easily distinguished from them by its shrubby perennial habit and oblong to ovate leaves with round tips (Flora of North America Editorial Committee, 2014). Crompton and Bassett (1985) provided a key that separates the closely related species, S. paulsenii, S. kali, S. collina and S. pestifer. 

Prevention and Control

Borger et al. (2009a,b) detailed the chemical, physical and biological control of S. australis [S. kali subsp. tragus]. See also Crompton and Bassett (1985), discussing S. kali subsp. ruthenica as S. pestifer A. Nels.

S. kali is a declared aquatic or terrestrial noxious weed and/or noxious-weed seed in several US states, including Arizona and Ohio as S. kali var. tenuifolia and California and Missouri as S. pestifer (USDA-ARS, 2015). As is common with this species complex, the taxonomy requires clarification.

Cultural control and sanitary measures

A two year study assessed five strategies for controlling S. tragus (here accepted as S. kali; see Notes of Taxonomy and Nomenclature) in Canada (Antill et al., 2012): removal of ungulate grazing; spraying with metsulfuron-methyl; manual pulling; broadcast seeding of native plant species, and integrated weed management combining grazing exclusion, herbicide and seeding. All treatments that included herbicide reduced S. tragus but also removed native forbs. Manual removal reduced weed cover and density but not final biomass, and was considered impractical to implement in large areas. Grazing exclusion resulted in a large decline of S. tragus density and biomass after two years and coincided with recovery of other forbs, suggesting that S. tragus responds to competition and could, therefore, be reduced through management of ungulate grazing pressure (Antill et al., 2012). Matooka et al. (2003) noted in Hawaii that S. kali is grazed, and fences stop the movement of detached plants, and both of which may be reasons why the species has not spread faster than it could have.

Physical/mechanical control

Young plants can be pulled or uprooted, or hoed just below ground level before seed set. Flowers can also be cut before maturity. Mowing tends to cause S. kali to adopt a low habit, but repeated mowing may provde control (PIER, 2015).

In all cases, it is important to establish desirable plants, such as competitive perennial grasses, in disturbed or open areas after any form of control to reduce re-invasion.

Cultivation, however, may promote growth of the weed. A study by Johnson and Fulbright (2008) found S. iberica on fine sandy loams to be 221 times greater on disked plots than on control plots. The authors stressed that land managers should consider soil series when disking for wildlife management, as disking disturbance can clearly worsen invasiveness on certain soils (Johnson and Fulbright, 2008).

Movement control

Attention should be taken regarding the movement of hay from areas where infestation is known. It is uncertain whether seeds survive the passage through livestock guts and therefore whether any quarantine for animals movements is warranted.

Biological control

The gall midge Desertovellum stackelbergi (Diptera: Cecidomyiidae), which attacks S. tragus in Uzbekistan, was identified as a candidate biological control agent for S. tragus in the USA (Sobhian et al., 2003). In a field test conducted in Uzbekistan, Californian S. tragus biotype A, similar to accessions from Uzbekistan, Greece and Ukraine, was the referred host, whereas Californian biotype B was distinct, though also attacked. Since both California biotypes were hosts to the gall midge, further studies on the biology and host specificity of the insect were considered justified (Sobhian et al., 2003).

The obligate biotrophic rust fungus Uromyces salsolae is a candidate biological control agent of S. tragus. Host range tests were conducted in quarantine and only species in the genus Salsola were susceptible to the fungus (Berner et al., 2009).

The eriophyid mite Aceria salsolae was also evaluated as a prospective classical biological control agent of invasive alien tumbleweeds, including Salsola tragus, S. collina, S. paulsenii and S. australis, in North America (Smith et al., 2009). S. tragus was heavily infested by A. salsolae and plant size was negatively correlated to the level of infestation. Although S. kali plants were also infested, plant size did not appear to be affected by the mites. Other nontarget plants were not as suitable for the mite in the field as in previous laboratory experiments, and it was concluded that there would be no significant risk to using A. salsolae as a biological agent in North America (Smith et al., 2009).

Two species of coleophorid moths are also reported to have been introduced to control S. tragus in western USA, where they became widely established; however, they were heavily attacked by predators and parasitoids and have not reduced S. tragus populations.

Cosmobaris scolopacea is a weevil distributed in Eurasia and North America, generally associated with the Chenopodiaceae family of flowering plants. The larvae feed and pupate within stems. Preliminary host range testing in Italy confirmed the presence of a highly divergent Sicilian lineage of C. scolopacea that was reared only on S. kali, indicating that this type requires further testing as a biological control agent (Cristofaro et al., 2011).

Colletotrichum gloeosporioides f. sp. salsolae is a facultative parasitic fungus being evaluated as a classical biological control agent of S. tragus in initial host range determination tests (Berner et al., 2012). Greenhouse tests showed that it is specific to Salsola spp., and that pathogen and inoculation procedures offer a low-cost solution to S. tragus infestations (Berner et al., 2014).

Despite the large number of identified potential biological control agents and host specificity testing, there appear to be few records of actual releases or more widespread field testing of the above.

Chemical control

Glyphosate can control S. kali if applied before seed set, whereas 2,4-D may actually cause S. kali to become tough and leathery, producing a plant that is more difficult to manage. It has also been observed that some S. tragus are showing resistance to sulfonylurea and trazine herbicides in western USA (PIER, 2015), though it is noted as very sensitive to hormone-type dicot herbicides (Motooka et al., 2003).

In northwestern USA, where it is a severe postharvest problem, S. tragus [Salsola kali subsp. tragus] is controlled either by tillage or applications of various herbicides. Paraquat plus diuron gave effective (>90%) control, whereas glyphosate plus 2,4-D provided unacceptable (≤75%) weed control. The use of laser controlled application led to a 42% reduction in chemical use compared with broadcast applications with a corresponding cost (Young et al., 2008).

In cotton fields, application of 2,4-D effectively controlled S. tragus, and could be applied close to the plant without injuring cotton, whereas dicamba and diflufenzopyr plus dicamba were less effective as size of the plant increased and both injured cotton regardless of interval between application and planting (Everitt and Keeling, 2007). Many studies have looked at various herbicide treatments and the economics of timing, application rate, and impacts on crop yields, including in sunflower (Godar, 2013) and beans (Arnold et al., 2012).

IPM

Borger et al. (2009b) concluded that effective weed management depended on: reducing seed dispersal; burning all senesced, mobile plants in late autumn, and herbicide control of the largest cohorts of S. australis in summer and autumn. Together, this reduced population growth rate. Using a model, they predicted that this control strategy resulted in a 66% chance of the population becoming extinct over 25 years.

Gaps in Knowledge/Research Needs

Salsola section kali requires a detailed literature review and thorough investigation, beginning with a full clarification of taxonomic limits of defined taxa in the native range and, where introduced, corresponding populations and the elucidation of parents in the case of hybrid populations. Only when this has been achieved and agreed can further advances be made on control. It is evident that different taxa have quite different environmental requirements and respond quite differently to different treatments. A full review of control methods, especially chemical and biological control, appears warranted.

References

Akhani H; Edwards G; Roalson EH, 2007. Diversification of the old world Salsoleae s.l. (Chenopodiaceae): molecular phylogenetic analysis of nuclear and chloroplast data sets and a revised classification. International Journal of Plant Sciences, 168(6):931-956. http://www.journals.uchicago.edu/IJPS/home.html

Akhani H; Greuter W; Roalson EH, 2014. Notes on the typification and nomenclature of Salsola and Kali (Chenopodiaceae). Taxon, 63(3):647-650. http://www.ingentaconnect.com/content/iapt/tax/2014/00000063/00000003/art00013

Antill TM; Naeth MA; Bork EW; Westhaver AL, 2012. Russian thistle (Salsola tragus L.) control on bighorn sheep winter ranges in Jasper National Park. Natural Areas Journal, 32(4):391-397. http://www.bioone.org/doi/abs/10.3375/043.032.0407

Arnold RN; Smeal D; Lombard KA; O'Neill MK; Allen SC; West M; Yazzie R, 2012. Broadleaf weed control in dry beans (Phaseolus vulgaris) with preemergence applications of valor alone or in combination from 2004 to 2008. Bulletin - Agricultural Experiment Station, New Mexico, No.804:4 pp. http://www.cahe.nmsu.edu/

Ayres D; Ryan FJ; Grotkopp E; Bailey J; Gaskin J, 2009. Tumbleweed (Salsola, section Kali) species and speciation in California. Biological Invasions, 11(5):1175-1187. http://www.springerlink.com/content/q785758119m43062/?p=4401958f27c2470391b4cef788abbd0b&pi=8

Beatley JC, 1973. Russian thistle (Salsola) species in western United States. Journal of Range Management, 26(3):225-226.

Beckie HJ; Francis A, 2009. The biology of Canadian weeds. 65. Salsola tragus L. (updated). Canadian Journal of Plant Science, 89(4):775-789. http://article.pubs.nrc-cnrc.gc.ca/RPAS/rpv?hm=HInit&calyLang=eng&journal=cjps&volume=89&afpf=CJPS08181.pdf

Berner D; Lagopodi AL; Kashefi J; Mukhina Z; Kolomiets T; Pankratova L; Kassanelly D; Cavin C; Smallwood E, 2014. Field assessment, in Greece and Russia, of the facultative saprophytic fungus, Colletotrichum salsolae, for biological control of Russian thistle (Salsola tragus). Biological Control, 76:114-123. http://www.sciencedirect.com/science/article/pii/S1049964414001248

Berner DK; Bruckart WL; Cavin CA; Michael JL, 2009. Mixed model analysis combining disease ratings and DNA sequences to determine host range of Uromyces salsolae for biological control of Russian thistle. Biological Control, 49(1):68-76. http://www.sciencedirect.com/science/journal/10499644

Berner DK; Cavin CA, 2012. Finalizing host range determination of a weed biological control pathogen with best linear unbiased predictors and damage assessment. BioControl [Proceedings of the "Biological Control for Nature" meeting, Northampton, Massachusetts, USA, 3-7 October 2010.], 57(2):235-246. http://www.springerlink.com/link.asp?id=102853

Berner DK; Cavin CA; McMahon MB; Loumbourdis I, 2006. First report of anthracnose of Salsola tragus caused by Colletotrichum gloeosporioides in Greece. Plant Disease, 90(7):971.

Blank RR; Young JA; Allen FL, 1999. Aeolian dust in a saline playa environment, Nevada, U.S.A. Journal of Arid Environments, 41(4):365-381.

Borger CPD; Scott JK, 2009. The biology of Australian weeds 55. Salsola australis R.Br. Plant Protection Quarterly, 24(4):126-137.

Borger CPD; Scott JK; Renton M; Walsh M; Powles SB, 2009. Assessment of management options for Salsola australis in south-west Australia by transition matrix modelling. Weed Research (Oxford), 49(4):400-408. http://www.blackwell-synergy.com/loi/wre

Borger CPD; Scott JK; Walsh M; Powles SB, 2009. Demography of Salsola australis populations in the agricultural region of south-west Australia. Weed Research (Oxford), 49(4):391-399. http://www.blackwell-synergy.com/loi/wre

Bruckart W; Cavin C; Vajna L; Schwarczinger I; Ryan FJ, 2004. Differential susceptibility of Russian thistle accessions to Colletotrichum gloeosporioides. Biological Control, 30(2):306-311.

CHAH (Council of Heads of Australasian Herbaria), 2015. Australia's virtual herbarium. Australia: Council of Heads of Australasian Herbaria. http://avh.ala.org.au

Cristofaro M; Lecce F; Paolini A; Cristina Fdi; Bon MC; Colonnelli E; Smith L, 2013. Host specificity of an Italian population of Cosmobaris scolopacea (Coleoptera: Curculionidae), candidate for the biological control of Salsola tragus (Chenopodiaceae). In: Proceedings of the XIII International Symposium on Biological Control of Weeds, Waikoloa, Hawaii, USA, 11-16 September, 2011 [ed. by Wu, Y.\Johnson, T.\Sing, S.\Raghu, S.\Wheeler, G.\Pratt, P.\Warner, K.\Center, T.\Goolsby, J.\Reardon, R.]. Hilo, USA: USDA Forest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry, 20-25.

Crompton CW; Bassett IJ, 1985. The biology of Canadian weeds. 65. Salsola pestifer A. Nels. Canadian Journal of Plant Science, 65(2):379-388.

Duke JA, 1983. Handbook of Energy Crops. https://www.hort.purdue.edu/newcrop/duke_energy/dukeindex.html

Evans RA; Young JA, 1980. Establishment of barbwire Russian thistle in desert environments. Journal of Range Management, 33(3):169-173.

Evans RA; Young JA, 1982. Russian thistle and barbwire Russian thistle seed and seedbed ecology. Agricultural Research Results, ARS, USDA, No.ARR-W-25. 40pp.

Everitt JD; Keeling JW, 2007. Weed control and cotton (Gossypium hirsutum) response to preplant applications of dicamba, 2,4-D, and diflufenzopyr plus dicamba. Weed Technology, 21(2):506-510. http://wssa.allenpress.com/perlserv/?request=get-document&doi=10.1614%2FWT-06-124.1

Flora of North America Editorial Committee, 2014. Flora of North America North of Mexico. http://www.efloras.org/flora_page.aspx?flora_id=1

Fowler JJ; Hageman JH, 1979. Russian-thistle, a potential forage for arid lands. In: Arid lands plant resources: proceedings of the international arid lands conference on plant resources, Texas Tech University [ed. by J. R. Goodin\D. K. Northington]. Lubbock, Texas, USA: Texas Tech University, International Center for Arid and Semi-Arid Land Studies (ICASALS)., 430-443.

Gaskin JF; Ryan FJ; Hrusa GF; Londo JP, 2006. Genotype diversity of Salsola tragus and potential origins of a previously unidentified invasive Salsola from California and Arizona. Madroño, 53(3):244-251.

Godar AS; Stahlman PW; Dille JA, 2013. Efficacy of tribenuron alone and following preemergence herbicides in tribenuron-resistant sunflower. Crop Management, No.June:CM-2013-0621-01-RS. http://www.plantmanagementnetwork.org/cm/element/sum2.aspx?id=10655

Johnson MVV; Fulbright TE, 2008. Is exotic plant invasion enhanced by a traditional wildlife habitat management technique? Journal of Arid Environments, 72(10):1911-1917. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WH9-4SPSPK2-1&_user=6686535&_coverDate=10%2F31%2F2008&_rdoc=16&_fmt=high&_orig=browse&_srch=doc-info(%23toc%236845%232008%23999279989%23695520%23FLA%23display%23Volume)&_cdi=6845&_sort=d&_docanchor=&_ct=22&_version=1&_urlVersion=0&_userid=6686535&md5=2cfe6363038a91195adfb3dc17ae4850

Kairo M; Ali B; Cheesman O; Haysom K; Murphy S, 2003. Report to the Nature Conservancy. Curepe, Trinidad and Tobago: CAB International, 132 pp.

Kolomiets T; Mukhina Z; Matveeva T; Bogomaz D; Berner DK; Cavin CA; Castlebury LA, 2009. First report of stem canker of Salsola tragus caused by Diaporthe eres in Russia. Plant Disease, 93(1):110. HTTP://www.apsnet.org

Kolomiets T; Skatenok O; Alexandrova A; Mukhina Z; Matveeva T; Bogomaz D; Berner DK; Cavin CA, 2008. First report of anthracnose of Salsola tragus caused by Colletotrichum gloeosporioides in Russia. Plant Disease, 92(9):1366. HTTP://www.apsnet.org

Longland WS, 1995. Desert rodents in disturbed shrub communities and their effects on plant recruitment. In: General Technical Report - Intermountain Research Station, USDA Forest Service, No. INT-GTR-315. 209-215.

Missouri Botanical Garden, 2015. Flora of China Checklist. Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/Project/FC

Missouri Botanical Garden, 2015. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

Mosyakin SL; Rilke S; Freitag H, 2014. (2323) Proposal to conserve the name Salsola (Chenopodiaceae s.str.; Amaranthaceae sensu APG) with a conserved type. Taxon, 63(5):1134-1135. http://www.ingentaconnect.com/content/iapt/tax/2014/00000063/00000005/art00023

PIER, 2015. Pacific Islands Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

Pyankov VI; Voznesenskaya EV; Kuz'min AN; Ku MSB; Ganko E; Franceschi VR; Black CC Jr; Edwards GE, 2000. Occurrence of C₃ and C₄ photosynthesis in cotyledons and leaves of Salsola species (Chenopodiaceae). Photosynthesis Research, 63(1):69-84.

Royal Botanic Garden Edinburgh, 2015. Flora Europaea. Edinburgh, UK: Royal Botanic Garden Edinburgh. http://rbg-web2.rbge.org.uk/FE/fe.html

Ryan FJ; Ayres DR, 2000. Molecular markers indicate two cryptic, genetically divergent populations of Russian thistle (Salsola tragus) in California. Canadian Journal of Botany, 78(1):59-67.

Ryan FJ; Bell DE; Sobhian R, 2002. What's in a name? Diversity and origin of Russian thistle in California through DNA markers. In: California Conference on Biological Control III, University of California at Davis, USA, 15-16 August, 2002 [ed. by Hoddle, M. S.]. Berkeley, USA: Center for Biological Control, College of Natural Resources, University of California, 51-56.

Smith L, 2005. Host plant specificity and potential impact of Aceria salsolae (Acari: Eriophyidae), an agent proposed for biological control of Russian thistle (Salsola tragus). Biological Control, 34(1):83-92. http://www.sciencedirect.com/science/journal/10499644

Smith L; Cristofaro M; Lillo Ede; Monfreda R; Paolini A, 2009. Field assessment of host plant specificity and potential effectiveness of a prospective biological control agent, Aceria salsolae, of Russian thistle, Salsola tragus. Biological Control, 48(3):237-243. http://www.sciencedirect.com/science/journal/10499644

Smith L; Hrusa GF; Gaskin JF, 2013. How many species of Salsola tumbleweeds (Russian thistle) occur in the Western USA? In: Proceedings of the XIII International Symposium on Biological Control of Weeds, Waikoloa, Hawaii, USA, 11-16 September, 2011 [ed. by Wu, Y.\Johnson, T.\Sing, S.\Raghu, S.\Wheeler, G.\Pratt, P.\Warner, K.\Center, T.\Goolsby, J.\Reardon, R.]. Hilo, USA: USDA Forest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry, 177.

Sobhian R; Ryan FJ; Khamraev A; Pitcairn MJ; Bell DE, 2003. DNA phenotyping to find a natural enemy in Uzbekistan for California biotypes of Salsola tragus L. Biological Control, 28(2):222-228.

The Plant List, 2013. The Plant List: a working list of all plant species. Version 1.1. London, UK: Royal Botanic Gardens, Kew. http://www.theplantlist.org

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

USDA-NRCS, 2015. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Wang XiaoWei; Chang ShuiJing; Dilixiati; Li Hui; Huang JunHua, 2008. Chromosome numbers and karyotypes of genus Salsola in Xinjiang. Acta Botanica Boreali-Occidentalia Sinica, 28(1):65-71. http://xbzwxb.nwsuaf.edu.cn

Young FL; Yenish JP; Launchbaugh GK; McGrew LL; Alldredge JR, 2008. Postharvest control of Russian thistle (Salsola tragus) with a reduced herbicide applicator in the Pacific Northwest. Weed Technology, 22(1):156-159. http://wssa.allenpress.com/perlserv/?request=get-abstract&doi=10.1614%2FWT-07-096.1

Young JA; Evans RA, 1979. Barbwire Russian thistle seed germination. Journal of Range Management, 32(5):390-394.

Young JA; Evans RA, 1979. Germination ecology of barbwire Russian thistle. In: Abstracts of 1979 Meeting of the Weed Science Society of America. 84.

Zakharyeva OI, 1985. Chromosome numbers of some flowering plants from the Caucasus and Middle Asia. Bot. Zhurn. Bot. Zhurn. SSSR, 70(12):1699-1701.

Links to Websites

Principal Source

Draft datasheet under review

Contributors

30/04/15 Original text by:

Nick Pasiecznik, consultant, France

Distribution Maps